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SMALL TERRESTRIAL MAMMALS SORICOMORPHA, CHIROPTERA, RODENTIA FROM THE EARLY HOLOCENE LAYERS OF MALA TRIGLAVCA SW SLOVENIA MALI TERESTIČNI SESALCI SORICOMORPHA, CHIROPTERA, RODENTIA IZ ZGODNJEHOLOCENSKIH PLASTI MALE TRIGLAVCE JZ SLOVENIJA Borut TOŠKAN 1 Abstract UDC 903.4(497.4)”627”:569.3 Izvleček UDK 903.4(497.4)”627”:569.3 Borut Toškan: Small terrestrial mammals (Soricomorpha, Borut Toškan: Mali terestični sesalci (Soricomorpha, Chirop- Chiroptera, Rodentia) from the Early Holocene layers of Mala tera, Rodentia) iz zgodnjeholocenskih plasti Male Triglavce Triglavca (SW Slovenia) (JZ Slovenija) At least 132 specimens belonging to no less than 21 species V zgodnjeholocenskih sedimentih iz Boreala jame Mala Tri- of small terrestrial mammals from the Boreal were identi- glavca pri Divači so bili najdeni ostanki najmanj 132 prim- $ed within the $nds from the Early Holocene sediments from erkov malih sesalcev, ki pripadajo vsaj 21 vrstam: Crocidura Mala Triglavca (the Kras Plateau, SW Slovenia), namely Croc- suaveolens, Sorex alpinus / araneus, S. minutus, Talpa cf. euro- idura suaveolens, Sorex alpinus / araneus, S. minutus, Talpa cf. paea, Barbastella barbastellus, Sciurus vulgaris, Cricetulus mi- europaea, Barbastella barbastellus, Sciurus vulgaris, Cricetulus gratorius, Arvicola terrestris, Microtus agrestis / arvalis, M. sub- migratorius, Arvicola terrestris, Microtus agrestis / arvalis, M. terraneus / liectensteini, Chionomys nivalis, Myodes glareolus, subterraneus / liectensteini, Chionomys nivalis, Myodes glareo- Dinaromys bogdanovi, Glis glis, Muscardinus avellanarius and lus, Dinaromys bogdanovi, Glis glis, Muscardinus avellanarius Apodemus avicollis / sylvaticus / agrarius / uralensis. Tedan- and Apodemus avicollis / sylvaticus / agrarius / uralensis. %is je združbe malih sesalcev kažejo na mozaičen tip habitata z small mammal assemblage indicates that their habitat was pre- mešanimi, pretežno listopadnimi gozdovi in manjšimi travniki sumably a mosaic of a mixed, predominantly deciduous forest, ter kamnišči. Primerjava z alohronimi mikrofavnami s Krasa je dotted with small meadows and some rocky substrate. A com- nakazala hiter favnistični obrat v zadnjih 5,000 letih. parison of allochronous microfaunas from the Kras Plateau in- Ključne besede: mali sesalci, Slovenija, Boreal, paleookolje. dicated a rapid faunistic change in the last 5,000 years. Keywords : small mammals, Slovenia, Boreal, palaeoenviron- ment. INTRODUCTION %e Early Holocene small mammal fauna of Slovenia is not washed through sieves. %us, in spite of numerous poorly known. %is is due to a small number of excavated attempts to treat the excavated sites in an all-round man- sites, on the one hand, and to the $eldwork techniques ner, including faunal remains (Pohar 1984, 1986, 1990; used in the past that were far from meeting the modern Toškan & Dirjec 2004, 2006), teeth and bones of small standards, on the other. Concerning the latter, an impor- mammals were only rarely collected (e.g. Rakovec 1959, tant weak point of the $eld work methodology worth 1961; Pohar 1986, 1990) and never analyzed in details mentioning here is that the excavated sediments were (for exception see Toškan & Kryštufek 2004). In the case 1 Institute of Archaeology SRC SASA, Novi trg 2, P.O. Box 306, SI-1001, Ljubljana, Slovenia, e-mail: [email protected] Received/Prejeto: 20.05.2008 ACTA CARSOLOGICA 38/1, 117-133, POSTOJNA 2009 BORUT TOŠKAN of Mala Triglavca, however, the wet-sieving of the sedi- mal faunas of the Kras Plateau. %e remains elaborated ments yielded a relatively rich small mammal assemblage in this paper are those of Soricomorpha, Chiroptera and (NISP = 749), which provide a good opportunity to ob- Rodentia. tain a deeper insight into the Early Holocene small mam- SITE, MATERIAL AND METHODS Mala Triglavca (herein a'er: M. Triglavca) is a pro- sional scaling on the rectangular symmetric matrix of nounced (8 m wide, 6 m long, 1.8–0.8 m high) rock shel- the Jaccard’s coe>cients of similarity. For measuring ter in a small collapsed doline of the Divača karst (437 m the percentage of species whose “temporal” ranges end a.s.l; Fig. 1). It was $rst described and measured for ar- within a chronologically distinct small mammal assem- chaeological purposes by F. Leben (1988); he excavated there in 1979-1985, depositing the excavated sediments in the vicinity of the rock-shelter. On his initiative, I. Turk later wet-sieved 1,677 kg of the deposited sedi- ment of the so-called Mesolithic horizon (Turk, 2004a), revealing also the small mammal remains elaborated in this paper. For the sake of reliability of the stratigraphic setting of the studied assemblage it was decided to com- pletely remove all the recent sediment overlying the Me- solithic material before beginning to sample. %at the bulk of the sediments in Leben’s deposits presumably belong to the Mesolithic layer is con$rmed by archaeo- logical $nds (Leben 1988; Turk & Turk 2004). Based on the number (N = 286) and type of the excavated stone artifacts, the available archaeological material is to be ascribed to the Sauveterrian-Castelnovian complex (ca. 8,000–6,000 B.C.; Turk & Turk 2004), which cor- responds to the Boreal palaeoclimatic period. In view of the above mentioned removal of the recent sediment deposited over the material of the so-called Mesolithic horizon as well as the relatively high number of collected stone artifacts datable to the Sauveterrian-Castelnovian complex, the same chronological frame is to be assumed also for the microfauna discussed below. %e small mammal material consists of teeth, max- illas, mandibles and postcranial remains, whereby the latter were not subjected to further analysis. %e com- parative recent and subfossil material (collections of the Natural History Museum of Slovenia and the Regional Museum Koper) originates mainly from Slovenia and Fig. 1: Map of the Kras Plateau and the archaeological sites (dots) adjacent regions of the Balkan Peninsula. Quantitative mentioned in the text. comparisons among taxa were based on the Number of Identi$ed Specimens (NISP) and on the Minimum Number of Individuals (MNI); cf. Grayson (1984, 17 +) blage, the Index of Faunistic Change (IFC) was used for further details. Within the framework of the biomet- (cf. Hagmeier & Stults 1964). %e relative hospitality of ric analysis, the existence of statistically signi$cant het- (palaeo)habitats to micro mammals and the prevalence erogeneity among individual samples was determined of environmental conditions needed to support each using the Sche+é test (StatSo' Inc. 2001). Allochronous species was assessed by packing the presence-absence small mammal faunas were compared by multidimen- matrix to the state of maximum nestedness ( cf. Atmar 118 ACTA CARSOLOGICA 38/1 – 2009 SMALL TERRESTRIAL MAMMALS SORICOMORPHA, CHIROPTERA, RODENTIA FROM THE EARLY HOLOCENE LAYERS ... & Patterson 1993). Statistical calculations were run in ture of molars follows Van Der Meulen (1973; arvico- StatSo' 2001 - Statistica for Windows (version 6.0) and lins) and Niethammer & Krapp (1978 for murins and in Nestedness temperature calculator programme (Stat 1982 for cricetins). All dimensions are given in mil- So' 2008). limeters. Cheek-teeth are abbreviated as follows: P – pre- Subfossil microfaunal remains from M. Triglavca molars, M – molars; capital letters indicate the upper are kept at the Regional Museum Koper. teeth and small letters the lower ones. The nomencla- RESULTS AND DISCUSSION TAXONOMY to the alpine ( Sorex alpinus Schinz, 1837) or the com- %e examined sample of the cave deposits yielded a total mon shrew ( S. araneus Linnaeus, 1758). Both species number of 980 molars, mandibles and maxillas of small are already known from the Early Holocene of the Kras mammals; 749 of these allowed identi$cation at least to Plateau (Toškan & Kryštufek 2004, 116 f); the common the generic level. %e material belonged to not less than shrew is represented also in the Upper Pleistocene layers 132 specimens, ascribed to 21 species from six families. of the Caverna degli Orsi/Medvedja jama cave (Boschian %e share of rodents within the identi$able items exceeds 1999-2000, 60). 97 percent, while bats, on the other hand, are represented by only 4 specimens. All species in the sample are still ex- Family: Moles (Talpidae) tant and most still occur in Slovenia. %e only exceptions are the grey hamster (Cricetulus migratorius) , Martino’s Talpa cf. europaea Linneaus, 1758 vole (Dinaromys bogdanovi) and the herb $eld mouse (Apodemus uralensis) . Genus Talpa is represented by a small number of Taxonomy and nomenclature follow Wilson & isolated teeth, which did not allow a reliable distinction Reeder (2005). between the common mole Talpa europaea and the blind mole T. caeca Savi, 1822. In spite of the size di+erences Order: Shrews and moles (Soricomorpha) (with T. europaea being bigger), the ranges of the two Family: Shrews (Soricidae) overlap considerably. %e best distinction was achieved on the basis of m1 and m 2 dimensions. Since the length Crocidura suaveolens (Pallas, 1811) and breadth of the two available m1 from M. Triglavca lie within the range of the recent T. europaeus (thus ex- %e lesser white-toothed shrew is represented by ceeding the maximal values for recent T. caeca ), only the four specimens: two fragmented rostrums and two iso- presence
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